Evidence of a component Allee effect driven by predispersal seed predation in a plant (Pedicularis rex,Orobanchaceae)

A small or sparse population may suffer a reduction in fitness owing to Allee effects. Here, we explored effects of plant density on pollination, reproduction and predation in the alpine herb Pedicularis rex over two years. We did not detect a significant difference in the pollination rate or fecundity (fruit set and the initial seed set) before predation between sparse and dense patches in either year, indicating no pollination-driven Allee effect. However, dense patches experienced significantly fewer attacks by predispersal seed predators in both years, resulting in a significantly decreased realized fecundity (final seed set), suggesting a component Allee effect driven by predispersal seed predation. Predation-driven Allee effects have been predicted by many models and demonstrated for a range of animals, but there is scant evidence for such effects in plants. Our study provides strong evidence of a component Allee effect driven by predation in a plant species.     

A test for Allee effects in the self‐incompatible wasp‐pollinated milkweed Gomphocarpus physocarpus

It has been suggested that plants that are good colonizers will generally have either an ability to self‐fertilize or a generalist pollination system. This prediction is based on the idea that these reproductive traits should confer resistance to Allee effects in founder populations and was tested using Gomphocarpus physocarpus (Asclepiadoideae: Apocynaceae), a species native to South Africa that is invasive in other parts of the world. We found no significant relationships between the size of G. physocarpus populations and various measures of pollination success (pollen deposition, pollen removal and pollen transfer efficiency) and fruit set. A breeding system experiment showed that plants in a South African population are genetically self‐incompatible and thus obligate outcrossers. Outcrossing is further enhanced by mechanical reconfiguration of removed pollinaria before the pollinia can be deposited. Self‐pollination is reduced when such reconfiguration exceeds the average duration of pollinator visits to a plant. Observations suggest that a wide variety of wasp species in the genera Belonogaster and Polistes (Vespidae) are the primary pollinators. We conclude that efficient pollination of plants in small founding populations, resulting from their generalist wasp‐pollination system, contributes in part to the colonizing success of G. physocarpus. The presence of similar wasps in other parts of the world has evidently facilitated the expansion of the range of this milkweed.     

Evaluating the effects of pollinator‐mediated interactions using pollen transfer networks: evidence of widespread facilitation in south Andean plant communities

Information about the relative importance of competitive or facilitative pollinator‐mediated interactions in a multi‐species context is limited. We studied interspecific pollen transfer (IPT) networks to evaluate quantity and quality effects of pollinator sharing among plant species on three high‐Andean communities at 1600, 1800 and 2000 m a.s.l. To estimate the sign of the effects (positive, neutral or negative), the relation between conspecific and heterospecific pollen deposited on stigmas was analysed with GLMMs. Network analyses showed that communities were characterised by the presence of pollen hub‐donors and receptors. We inferred that facilitative and neutral pollinator‐mediated interactions among plants prevailed over competition. Thus, the benefits from pollinator sharing seem to outweigh the costs (i.e. heterospecific deposition and conspecific pollen loss). The largest proportion of facilitated species was found at the highest elevation community, suggesting that under unfavourable conditions for the pollination service and at lower plant densities facilitation can be more common.     

Experimental reduction of pollinator visitation modifies plant–plant interactions for pollination

The strength of interactions between plants for pollination depends on the abundance of plants and pollinators in the community. The abundance of pollinators may influence plant associations and densities at which individual fitness is maximized. Reduced pollinator visitation may therefore affect the way plant species interact for pollination. We experimentally reduced pollinator visitation to six pollinator‐dependent species (three from an alpine and three from a lowland community in Norway) to study how interactions for pollination were modified by reduced pollinator availability. We related flower visitation, pollen limitation and seed set to density of conspecifics and pollinator‐sharing heterospecifics inside 30 dome‐shaped cages partially covered with fishnet (experimental plots) and in 30 control plots. We expected to find stronger interactions between plants in experimental compared to controls plots. The experiment modified plant–plant interactions for pollination in all the six species; although for two of them neighbourhood interactions did not affect seed set. The pollen limitation and seed set data showed that reduction of pollinator visits most frequently resulted in novel and/or stronger interactions between plants in the experimental plots that did not occur in the controls. Although the responses were species‐specific, there was a tendency for increasing facilitative interactions with conspecific neighbours in experimental plots where pollinator availability was reduced. Heterospecifics only influenced pollination and fecundity in species from the alpine community and in the experimental plots, where they competed with the focal species for pollination. The patterns observed for visitation rates differed from those for fecundity, with more significant interactions between plants in the controls in both communities. This study warns against the exclusive use of visitation data to interpret plant–plant interactions for pollination, and helps to understand how plant aggregations may buffer or intensify the effects of a pollinator loss on plant fitness.     

Plant population dynamics, pollinator foraging, and the selection of self-fertilization

Many flowering plants rely on pollinators, self-fertilization, or both for reproduction. We model the consequences of these features for plant population dynamics and mating system evolution. Our mating systems-based population dynamics model includes an Allee effect. This often leads to an extinction threshold, defined as a density below which population densities decrease. Reliance on generalist pollinators who primarily visit higher density plant species increases the extinction threshold, whereas autonomous modes of selfing decrease and can eliminate the threshold. Generalist pollinators visiting higher density plant species coupled with autonomous selfing may introduce an effect where populations decreasing in density below the extinction threshold may nonetheless persist through selfing. The extinction threshold and selfing at low density result in populations where individuals adopting a single reproductive strategy exhibit mating systems that depend on population density. The ecological and evolutionary analyses provide a mechanism where prior selfing evolves even though inbreeding depression is greater than one-half. Simultaneous consideration of ecological and evolutionary dynamics confirms unusual features (e.g., evolution into extinction or abrupt increases in population density) implicit in our separate consideration of ecological and evolutionary scenarios. Our analysis has consequences for understanding pollen limitation, reproductive assurance, and the evolution of mating systems.     

Allee effects,extinctions, and chaotic transients in simple population models

Discrete time single species models with overcompensating density dependence and an Allee effect due to predator satiation and mating limitation are investigated. The models exhibit four behaviors: persistence for all initial population densities, bistability in which a population persists for intermediate initial densities and otherwise goes extinct, extinction for all initial densities, and essential extinction in which "almost every" initial density leads to extinction. For fast-growing populations, these models show populations can persist at high levels of predation even though lower levels of predation lead to essential extinction. Alternatively, increasing the predator's handling time, the population's carrying capacity, or the likelihood of mating success may lead to essential extinction. In each of these cases, the mechanism behind these disappearances are chaotic dynamics driving populations below a critical threshold determined by the Allee effect. These disappearances are proceeded by chaotic transients that are proven to be approximately exponentially distributed in length and highly sensitive to initial population densities.     

Modeling the Effect of Prey Refuge on a Ratio-Dependent Predator–Prey System with the Allee Effect

The extinction of species is a major threat to the biodiversity. The species exhibiting a strong Allee effect are vulnerable to extinction due to predation. The refuge used by species having a strong Allee effect may affect their predation and hence extinction risk. A mathematical study of such behavioral phenomenon may aid in management of many endangered species. However, a little attention has been paid in this direction. In this paper, we have studied the impact of a constant prey refuge on the dynamics of a ratio-dependent predator–prey system with strong Allee effect in prey growth. The stability analysis of the model has been carried out, and a comprehensive bifurcation analysis is presented. It is found that if prey refuge is less than the Allee threshold, the incorporation of prey refuge increases the threshold values of the predation rate and conversion efficiency at which unconditional extinction occurs. Moreover, if the prey refuge is greater than the Allee threshold, situation of unconditional extinction may not occur. It is found that at a critical value of prey refuge, which is greater than the Allee threshold but less than the carrying capacity of prey population, system undergoes cusp bifurcation and the rich spectrum of dynamics exhibited by the system disappears if the prey refuge is increased further.     

Mycorrhizal symbiosis increases the benefits of plant facilitative interactions

The diversity of pathways through which mycorrhizal fungi alter plant coexistence hinders the understanding of their effects on plant‐plant interactions. The outcome of plant facilitative interactions can be indirectly affected by mycorrhizal symbiosis, ultimately shaping biodiversity patterns. We tested whether mycorrhizal symbiosis enhances plant facilitative interactions and whether its effect is consistent across different methodological approaches and biological scenarios. We conducted a meta‐analysis of 215 cases (involving 21 nurse and 29 facilitated species), in which the performance of a facilitated plant species is measured in the presence or absence of mycorrhizal fungi. We show that mycorrhizal fungi significantly enhance plant facilitative interactions mainly through an increment in plant biomass (aboveground) and nutrient content, although their effects differ across biological contexts. In semiarid environments mycorrhizal symbiosis enhances plant facilitation, while its effect is non‐significant in temperate ecosystems. In addition, arbuscular but not ecto‐mycorrhizal (EMF) fungi significantly enhance plant facilitation, particularly increasing the P content of the plants more than EMF. Some knowledge gaps regarding the importance of this phenomenon have been detected in this meta‐analysis. The effect of mycorrhizal symbiosis on plant facilitation has rarely been assessed in other ecosystems different from semiarid and temperate forests, and rarely considering other fungal benefits provided to plants besides nutrients. Finally, we are still far from understanding the effects of the whole fungal community on plant‐plant interactions, and on plant species coexistence.     

Number of conspecifics and reproduction in the invasive plant Eschscholzia californica (Papaveraceae): is there a pollinator‐mediated Allee effect?

The component Allee effect has been defined as ‘a positive relationship between any measure of individual fitness and the number or density of conspecifics’. Larger plant populations or large patches have shown a higher pollinator visitation rate, which may give rise to an Allee effect in reproduction of the plants. We experimentally tested the effect of number of conspecifics on reproduction and pollinator visitation in Eschscholzia californica Cham., an invasive plant in Chile. We then built patches with two, eight and 16 flowering individuals of E. californica (11 replicates per treatment) in an area characterised by dominance of the study species. We found that E. californica exhibits a component Allee effect, as the number of individuals of this species has a positive effect on individual seed set. However, individual fruit production was not affected by the number of plants examined. Pollinator visitation rate was also independent of the number of plants, so this factor would not explain the Allee effect. This rate was positively correlated with the total number of flowers in the patches. We also found that the number of plants did not affect the seed mass or proportion of germinated seeds in the patches. Higher pollen availability in patches with 16 plants and pollination by wind could explain the Allee effect. The component Allee effect identified could lead to a weak demographic Allee effect that might reduce the rate of spread of E. californica. Knowledge of this would be useful for management of this invasive plant in Chile.     

Neutral hybridization can overcome a strong Allee effect by improving pollination quality

Small populations of plant species can be susceptible to demographic Allee effects mainly due to pollen limitation. Although sympatry with a common, co-flowering species may somewhat alleviate the problem of pollinator visitation (pollination quantity), the interspecific pollen transfer, IPT, (pollination quality) may remain a barrier to reproduction in small populations such as new introductions. However, if the two species are crosscompatible, our hypothesis is that neutral hybridization can help the small founding population overcome the Allee effect by improving the quality of pollination. We tested this hypothesis by using a novel modelling approach based on the theory of kinetic reactions wherein pollinators act as enzymes to catalyse the reaction between the two substrates: pollen and unselfed ovule. Using a single locus, two-allele genetic model, we developed a generic model that allows for hybridization between the invading and the native genotypes. Analysing the stability properties of the trivial equilibria in hybridization model as compared with the single genotype invasion model, we found that hybridization can either remove or reduce the Allee effect by making an otherwise stable trivial equilibrium unstable. Our study suggests that hybridization can be neutral but still be the key driver of a successful invasion by mediating pollen limitation. Conservation programmes should therefore account for this cryptic role that hybridization could play in plant invasions.     

How to differentiate facilitation and environmentally driven co‐existence

Positive plant–plant interactions (i.e. facilitation) receive increasing attention as a potentially important driver of community assembly. We conducted a systematic literature review indicating broad support for positive effects of potential facilitator species. However, a large majority of the reviewed studies (83% for field studies, 57% for experiments) share a similar risk of misinterpretation as they assess facilitative effects by comparing plots inhabited by a potential facilitator with randomly placed control plots nearby (paired sampling). As the distribution of facilitator species may itself be environmentally driven, species co‐existence caused by facilitation cannot exclusively be separated from environmental effects (habitat sharing). Based on simulated plant communities and sampling protocols, we show how non‐random co‐existence can occur in the absence of facilitation. This is relevant because both the effect of spatial environmental heterogeneity and of facilitation (stress‐gradient hypothesis) are expected to increase with environmental harshness. Nevertheless, 58% of facilitation studies neither undertook measures to minimize potential biases in their sampling approaches nor did they acknowledge such limitations in the discussion. Attention to this problem has significantly decreased in recent years. We propose that facilitation studies could be improved by (1) using random sampling for association studies, (2) co‐analysing environmental factors, or (3) experimentally establishing presumed facilitators. Experimental approaches mimicking facilitative plant characteristics can help to identify facilitation mechanisms. Combining approaches and including functional traits of the involved species should further strengthen inference.     

Comparison of the ecology and evolution of plants with a generalist bird pollination system between continents and islands worldwide

Thousands of plant species worldwide are dependent on birds for pollination. While the ecology and evolution of interactions between specialist nectarivorous birds and the plants they pollinate is relatively well understood, very little is known on pollination by generalist birds. The flower characters of this pollination syndrome are clearly defined but the geographical distribution patterns, habitat preferences and ecological factors driving the evolution of generalist‐bird‐pollinated plant species have never been analysed. Herein I provide an overview, compare the distribution of character states for plants growing on continents with those occurring on oceanic islands and discuss the environmental factors driving the evolution of both groups. The ecological niches of generalist‐bird‐pollinated plant species differ: on continents these plants mainly occur in habitats with pronounced climatic seasonality whereas on islands generalist‐bird‐pollinated plant species mainly occur in evergreen forests. Further, on continents generalist‐bird‐pollinated plant species are mostly shrubs and other large woody species producing numerous flowers with a self‐incompatible reproductive system, while on islands they are mostly small shrubs producing fewer flowers and are self‐compatible. This difference in character states indicates that diverging ecological factors are likely to have driven the evolution of these groups: on continents, plants that evolved generalist bird pollination escape from pollinator groups that tend to maintain self‐pollination by installing feeding territories in single flowering trees or shrubs, such as social bees or specialist nectarivorous birds. This pattern is more pronounced in the New compared to the Old World. By contrast, on islands, plants evolved generalist bird pollination as an adaptation to birds as a reliable pollinator group, a pattern previously known from plants pollinated by specialist nectarivorous birds in tropical mountain ranges. Additionally, I discuss the evolutionary origins of bird pollination systems in comparison to systems involving specialist nectarivorous birds and reconstruct the bird pollination system of Hawaii, which may represent an intermediate between a specialist and generalist bird pollination system. I also discuss the interesting case of Australia, where it is difficult to distinguish between specialist and generalist bird pollination systems.     

Plant sex alters Allee effects in aggregating plant parasites

Species interactions are central to our understanding of population dynamics. While density typically strengthens competition, reducing absolute fitness, Allee effects can reverse this pattern, increasing fitness with density. Allee effects emerge in host–parasite systems when higher parasite densities dilute immune responses or increase resource‐mobilization. The optimal density of individuals in these systems should be influenced by how host quality alters the rates at which facilitative and competitive effects change across densities. We tested these ideas using sumac Rhus typhina and a gall‐forming parasite Melaphis rhois that attacks sumac leaves. Fitness peaked at intermediate densities, indicating an Allee effect, but the fitness peak depended on host sex. Patterns of abundance mirrored fitness patterns, with galls clustered on leaves and female hosts supporting greater numbers of galls. Within leaves, galls near the stem were more fit, and gall‐makers preferentially oviposited near to the stem. The patterns of fitness and abundance are consistent with Allee effects caused by increased resource mobilization at higher gall‐maker densities rather than diluted immune responses. Our results suggest that Allee effects in parasites can be described as the summative effects of competitive and facilitative processes and, because both are common, Allee effects are likely common in host–parasite systems.     

开花时间与伴生种对鹤首马先蒿传粉和生殖成功的影响

开花物候是物种间相互作用的重要生活史特征和适合度因子,在全球气候变化的背景下而备受关注.为探讨开花时间如何存种内和种间水平上影响植物的传粉和生殖成功,我们连续3年(2003-2005)对不同花期和伴生种存在情况下的鹤首马先蒿(Pedicularis gruina)的传粉者访花忠实度、受粉率、坐果率、单果种子产量和果实被啃食频率进行了比较研究.结果表明鹤首马先蒿的坐果率主要受其传粉环境的影响:在没有伴生种时,不同时期鹤首马先蒿的坐果率没有显著差异,34-38％的花可以坐果;在有伴生种存在时既可以显著提高其坐果率,也可显著降低其坐果率,这取决于传粉者类型以及伴生种密穗马先蒿(P.densispica)花期的差异.密穗马先蒿具有花蜜和花粉双重报酬,在群落中可以作为主体物种吸引传粉者并间接促进与其伴生的鹤首马先蒿的传粉和生殖成功.同样无蜜的管花马先蒿(P.siphonantha)和鹤首马先蒿伴生,则是通过提高群落水平对传粉者的吸引力进而直接促进鹤首马先蒿的传粉和生殖成功.此外,研究结果也表明开花时间对坐果率没有显著影响,但是显著影响单果种子产量和果实被啃食的频率.在相同的传粉条件下早花期植株单果种子产量显著高于晚花期的种子产量,同时早花期的果实被啃食的频率显著增加.     

The nested assembly of plant facilitation networks prevents species extinctions

Facilitation is a positive interaction assembling ecological communities and preserving global biodiversity. Although communities acquire emerging properties when many species interact, most of our knowledge about facilitation is based on studies between pairs of species. To understand how plant facilitation preserves biodiversity in complex ecological communities, we propose to move from the study of pairwise interactions to the network approach. We show that facilitation networks behave as mutualistic networks do, characterized by a nonrandom, nested structure of plant-plant interactions in which a few generalist nurses facilitate a large number of species while the rest of the nurses facilitate only a subset of them. Consequently, generalist nurses shape a dense and highly connected network. Interestingly, such generalist nurses are the most abundant species in the community, making facilitation-shaped communities strongly resistant to extinction, as revealed by coextinction simulations. The nested structure of facilitative networks explains why facilitation, by preventing extinction, preserves biodiversity.     

When can two plant species facilitate each other's pollination?

Facilitation occurs when an increase in the density of one species causes an increase in the population growth rate or the density of a second species. In plants, ample evidence demonstrates that one species can facilitate another by ameliorating abiotic conditions, but the hypothesis that pollination facilitation – in which the presence of one flowering species increases pollinator visits to a second species – can also occur remains controversial. To identify the necessary conditions for pollination facilitation to occur, we constructed population models of two plant species that share the same pollinator and compete for establishment sites, and we assumed that heterospecific pollen can interfere with successful seed set. We found that facilitation for pollination occurs only when the pollinator visitation rate is an initially accelerating function of the combined numbers of flowering plants of both species in a patch. The presence of a second species can allow populations of a focal species either to persist for a longer amount of time before going extinct ("weak facilitation") or to persist indefinitely at a stable equilibrium density ("strong facilitation"). When only a single plant of either species can occupy a site, the plant species with the higher initial density can experience strong facilitation but will eventually out-compete the other species. However, when site occupancy was not exclusive, strong facilitation sometimes led to coexistence of the two species. Increasing the extent of pollen carryover increased the range of initial population densities leading to strong facilitation. In light of our theoretical results, we discuss the apparent rarity of pollination facilitation in nature.     

Reproductive success in fragmented habitats: do compatibility systems and pollination specialization matter?

Abstract. This paper explores whether plant breeding system and pollination specialization influence the reproductive response of plants to habitat fragmentation. It is meaningful for conservation to predict a plant species’ extinction risk. We found 25 studies in the literature assessing the effects of habitat fragmentation on either pollination or reproductive success of 46 plant species to answer the following questions: 1. Are pollination and reproductive success of self‐incompatible species more likely to decline with habitat fragmentation than the pollination and reproductive success of self‐compatible species? Although most of the species showed statistically significant negative effects, the pollination and reproduction of self‐incompatible species were as likely to decline with fragmentation as those of self‐compatible species. 2. Are pollination and reproductive success of specialist plants more affected than the pollination and reproduction of generalist plants? Comparisons of fragmentation‐related changes in pollination and reproductive success between specialists and generalists do not support the hypothesis that specialization in pollination increases the risk of plant extinction. 3. Can self‐incompatible species offset their expected higher vulnerability to fragmentation by being, on average, more pollination generalist than self‐compatible species? In a larger data set on 260 species, we did not find significant differences in either the mean number or frequency distribution of numbers of flower‐visiting species or orders between self‐compatible and self‐incompatible species. Our review suggests that no generalizations can be made on susceptibility to fragmentation based on compatibility system and pollination specialization.     

Impacts of predation on dynamics of age-structured prey: Allee effects and multi-stability

With a series of mathematical models, we explore impacts of predation on a prey population structured into two age classes, juveniles and adults, assuming generalist, age-specific predators. Predation on any age class is either absent, or represented by types II or III functional responses, in various combinations. We look for Allee effects or more generally for multiple stable steady states in the prey population. One of our key findings is the occurrence of a predator pit (low-density ??refuge?? state of prey induced by predation; the chance of escaping predation thus increases both below and above an intermediate prey density) when only one age class is consumed and predators use a type II functional response ??this scenario is known to occur for an unstructured prey consumed via a type III functional response and can never occur for an unstructured prey consumed via a type II one. In the case where both age classes are consumed by type II generalist predators, an Allee effect occurs frequently, but some parameters give also rise to a predator pit and even three stable equilibria (one extinction equilibrium and two positive ones??Allee effect and predator pit combined). Multiple positive stable equilibria are common if one age class is consumed via a type II functional response and the other via a type III functional response??here, in addition to the behaviours mentioned above one may even observe three stable positive equilibria????double?? predator pit. Some of these results are discussed from the perspective of population management.     

The role of Allee effects in gypsy moth, <Emphasis Type="Italic">Lymantria dispar</Emphasis> (L.), invasions

Allee effects have been applied historically in efforts to understand the low-density population dynamics of rare and endangered species. Many biological invasions likewise experience the phenomenon of decreasing population growth rates at low population densities because most founding populations of introduced nonnative species occur at low densities. In range expansion of established species, the initial colonizers of habitat beyond the organism’s current range are usually at low density, and thus could be subject to Allee dynamics. There has been consistent empirical and theoretical evidence demonstrating, and in some cases quantifying, the role of Allee dynamics in the gypsy moth, Lymantria dispar (L.), invasion of North America. In this review, we examine the potential causes of the Allee effect in the gypsy moth and highlight the importance of mate-finding failure as a primary mechanism behind an Allee effect, while the degree to which generalist predators induce an Allee effect remains unclear. We then explore the role of Allee effects in the establishment and spread dynamics of the gypsy moth system, which conceptually could serve as a model system for understanding how Allee effects manifest themselves in the dynamics of biological invasions.     

Pollinator coupling can induce synchronized flowering in different plant species

Synchronous and intermittent plant reproduction has been identified widely in diverse biomes. While synchronous flowering is normally observed within the same species, different species also flower in synchrony. A well-known example of interspecific synchrony is “general flowering" in tropical rain forests of Southeast Asia. Environmental factors, such as low temperature and drought, have been considered as major trigger of general flowering. However, environmental cues are not enough to explain general flowering because some trees do not flower even when they encounter favorable environmental cues. We propose alternative explanation of general flowering; “pollinator coupling”. When species flower synchronously, the elevated pollen and nectar resource may attract increased numbers of generalist pollinators, with a concomitant enhancement of pollination success (facilitation). However, under these circumstances, plants of different species may compete with one another for limited pollinator services, resulting in declines in pollination success for individual species (competition). Here, we present a model describing resource dynamics of individual trees serviced by generalist pollinators. We analyze combinations of conditions under which plants reproduce intermittently with synchronization within species, and/or (sometimes) between different species. We show that plants synchronize flowering when the number of pollinators attracted to an area increases at an accelerating rate with increasing numbers of flowers. In this case, facilitation of flowering by different species exceeds the negative influence of interspecific plant competition. We demonstrate mathematically that co-flowering of different species occurs under a much narrower range of circumstances than intraspecific co-flowering.